Leopard Cat (Prionailurus bengalensis)
The leopard cat (Prionailurus bengalensis) is a small
wild cat of South and East Asia. Since 2002 it has been listed as Least Concern
by IUCN as it is widely distributed but threatened by habitat loss and hunting
in parts of its range. There are twelve leopard cat subspecies, which differ
widely in appearance. The leopard cat's name is derived from the leopard-like
spots prevalent in all subspecies, but its relation to the leopard is distant.
Leopard cats are about the size of a domestic cat, but more slender with longer
legs and well-defined webs between the toes. Their small head is marked with two
prominent dark stripes, their short and narrow muzzle white. There are two dark
stripes running from the eyes to the ears, and smaller white streaks running
from the eyes to the nose. The backs of their moderately long and rounded ears
are black with a central white spot. Body and limbs are marked with black spots
of varying size and colour, and along the back are two to four rows of elongated
spots. The tail is about half the size of their head-body-length and spotted
with a few indistinct rings near the black tip. The background colour of their
spotted fur is tawny with a white chest and belly. But in their huge range, they
vary so much in coloration and size of spots as well as in body size and weight
that initially they were thought to be several different species. The fur colour
is yellowish brown in the southern populations, but pale silver-grey in the
northern ones. The black markings may be spotted, rosetted, or even forming
dotted streaks, depending on the subspecies. In the tropics, leopard cats weigh
0.55 to 3.8 kg (1.2 to 8.4 lb), have a head-body-length of 38.8 to 66 cm (15.3
to 26 in) with a 17.2 to 31 cm (6.8 to 12 in) long tail. In northern China and
Siberia, they weigh up to 7.1 kg (16 lb), and have a head-body-length of up to
75 cm (30 in); generally, they put on weight before winter and become thinner
until spring. Shoulder height is about 41 cm (16 in). In 1792, Robert Kerr first
described a leopard cat under the binominal Felis bengalensis in his
translation of Carl von Linné’s Systema Naturae as being native to southern
Bengal. Between 1829 and 1922, different authors of 20 more descriptions
classified the cat either as Felis or Leopardus.[ Owing to the individual
variation in fur colour, leopard cats from British India were described as
Felis nipalensis from Nepal, Leopardus ellioti from the area of Bombay,
Felis wagati and Felis tenasserimensis from Tenasserim. In 1939,
Reginald Innes Pocock subordinated them to the genus Prionailurus. The
collection of the Natural History Museum in London comprised several skulls and
large amounts of skins of leopard cats from various regions. Based on this broad
variety of skins, he proposed to differentiate between a southern subspecies
Prionailurus bengalensis bengalensis from warmer latitudes to the west and
east of the Bay of Bengal, and a northern Prionailurus bengalensis horsfieldi
from the Himalayas, having a fuller winter coat than the southern. His
description of leopard cats from the areas of Gilgit and Karachi under the
trinomen Prionailurus bengalensis trevelyani is based on seven skins that
had longer, paler and more greyish fur than those from the Himalayas. He assumed
that trevelyani inhabits more rocky, less forested habitats than bengalensis and
horsfieldi. Between 1837 and 1930, skins and skulls from China were described as
Felis chinensis, Leopardus reevesii, Felis scripta, Felis microtis,
decolorata, ricketti, ingrami, anastasiae and sinensis, and later grouped
under the trinomen Felis bengalensis chinensis.In the beginning of the
20th century, a British explorer collected wild cat skins on the island of
Tsushima. Oldfield Thomas classified these as Felis microtis, which had
been first described by Henri Milne-Edwards in 1872. Two skins from Siberia
motivated Daniel Giraud Elliot to write a detailed description of Felis
euptilura in 1871. One was depicted in Gustav Radde’s illustration cum
description of a wild cat; the other was part of a collection at the Regent's
Park Zoo. The ground colour of both was light brownish-yellow, strongly mixed
with grey and covered with reddish-brown spots, head grey with a dark-red stripe
across the cheek. In 1922, Tamezo Mori described a similar but lighter grey
spotted skin of a wild cat from the vicinity of Mukden in Manchuria that he
named Felis manchurica. Later both were grouped under the trinomen
Felis bengalensis euptilura. In the 1970s and 1980s, the Russian zoologists
Geptner, Gromov and Baranova disagreed with this classification. They emphasized
the differences of skins and skulls at their disposal and the ones originating
in Southeast Asia, and coined the term Amur forest cat, which they regarded as a
distinct species. In 1987, Chinese zoologists pointed out the affinity of
leopard cats from northern China, Amur cats and leopard cats from southern
latitudes. In view of the morphological similarities they did not support
classifying the Amur cat as a species.
The initial binomial euptilura given by Elliott was eventually changed to euptilurus referring to the ICZN Principle of Gender Agreement; at present, both terms are used.
Distribution and Habitat
Leopard cats are the most widely distributed Asian small cats. Their range extends from the Amur region in the Russian Far East over the Korean Peninsula, China, Indochina, the Indian Subcontinent, to the West in northern Pakistan, and to the south in the Philippines and the Sunda islands of Indonesia. They are found in agriculturally used areas but prefer forested habitats. They live in tropical evergreen rainforests and plantations at sea level, in subtropical deciduous and coniferous forests in the foothills of the Himalayas at altitudes above 1,000 m (3,300 ft). In 2009, a leopard cat was camera trapped in Nepal’s Makalu-Barun National Park at an altitude of 3,254 m (10,676 ft). At least six individuals inhabit the survey area, which is dominated by associations of rhododendron, oak and maple. In the northeast of their range they live close to rivers, valleys and in ravine forests, but avoid areas with more than 10 cm (3.9 in) of snowfall. They are rare in Pakistan’s arid treeless areas.
In Sabah’s Tabin Wildlife Reserve leopard cats had average home ranges of 3.5 km2 (1.4 sq mi). In Thailand’s Phu Khieu Wildlife Reserve 20 leopard cats were radio-collared between 1999 and 2003. Home ranges of males ranged from 2.2 km2 (0.85 sq mi) to 28.9 km2 (11.2 sq mi), and of the six females from 4.4 km2 (1.7 sq mi) to 37.1 km2 (14.3 sq mi) The leopard cat is a widespread species in Asia. It is found throughout most of India west into Pakistan and Afghanistan (Habibi 2004), through the Himalayan foothills, across most of China, and north to the Korean peninsula and into the Russian Far East (Nowell and Jackson 1996). It is found throughout Southeast Asia, and on the islands of Sumatra, Java, Borneo and Taiwan. It is found on numerous small offshore islands of mainland Asia (Nowell and Jackson 1996, Sunquist and Sunquist 2002). The leopard cat is the only wild felid found in the Japan, where it occurs on the small islands Tsushima and Iriomote, and the Philippines, where it occurs on the islands of Palawan, Panay, Negros and Cebu. In the Philippines, there are recent (2007) unconfirmed reports from the island of Masbate. It should be present in Guimaras due to proximity to Negros and Panay, but no presence was reported, and is therefore presumed to be extinct (R. Lorica and W. Oliver, unpub.).
Afghanistan; Bangladesh; Bhutan; Brunei Darussalam; Cambodia; China; Hong Kong; India; Indonesia (Jawa, Kalimantan, Sumatera); Japan (Nansei-shoto); Korea, Democratic People's Republic of; Korea, Republic of; Lao People's Democratic Republic; Malaysia; Myanmar; Nepal; Pakistan; Philippines; Russian Federation; Singapore; Taiwan, Province of China; Thailand; Viet Nam
Distribution of subspecies
As of 2009, the following subspecies are recognized:
Prionailurus bengalensis bengalensis (Kerr, 1792) — India, Bangladesh, Myanmar, Thailand, the Malay Peninsula, Indochina to Yunnan in China.
Prionailurus bengalensis javanensis (Desmarest, 1816) — Java and Bali.
Prionailurus bengalensis sumatranus (Horsfield 1821) — Sumatra and Tebingtinggi.
Prionailurus bengalensis chinensis (Gray 1837) — Taiwan and China except Yunnan.
Prionailurus bengalensis horsfieldi (Gray 1842) — Kashmir, Punjab, Kumaon, Nepal, Sikkim and Bhutan.
Prionailurus bengalensis euptilurus/euptilura, the Amur cat, (Elliott 1871) — eastern Siberia, in Manchuria, in Korea and on the Tsushima Island in the Korea Strait.
Prionailurus bengalensis borneoensis (Brongersma 1936) — Borneo.
Prionailurus bengalensis trevelyani (Pocock 1939) — northern Kashmir and Punjab, and in Southern Baluchistan.
Prionailurus bengalensis alleni (Sody, 1949) — Hainan Island.
Prionailurus bengalensis iriomotensis (Imaizumi, 1967) — found exclusively on the tiny island of Iriomote, one of the Ryukyu Islands in the Japanese Archipelago.
Prionailurus bengalensis heaneyi (Groves 1997) — the Philippine island of Palawan.
Prionailurus bengalensis rabori (Groves 1997) — the Philippine islands of Negros, Cebu, and Panay.
The Iriomote cat (P.b. iriomotensis) has been proposed as a species since 1967, but following mtDNA analysis in the 1990s is now considered a subspecies of the leopard cat.
The Tsushima leopard cat lives exclusively on Tsushima Island. Initially regarded as belonging to the Chinese leopard cat subspecies, it is now considered an isolated population of the Amur cat P. b. euptilurus/euptilura
The leopard cat is the most frequently recorded small cat
across most of its wide range, in comparison with sympatric species (Nowell and
Jackson 1996, Duckworth et al. 1999, Holden 2001, Duckworth et al. 2005, Lynam
et al. 2006, Yasuda et al. 2007), and with its broad distribution has an
abundant population. However, it is probably declining due to habitat loss and
hunting. Large numbers of leopard cat furs were exported from China (averaging
200,000 skins per year in the late 1980s) (Nowell and Jackson 1996). Higher
survival rates (92%) were recorded in a protected area with little human
influence, compared with lower rates in areas with greater human activity
(53-82%) (Haines et al. 2004). While the leopard cat is more tolerant of
disturbed areas than other small Asian felids, it likely undergoes higher
mortality in such areas. Island populations are most at risk of extinction, with
the Iriomote cat P.b. iriomotensis listed as Critically Endangered, and
the Visayan leopard cat P.b. rabori of the Philippine islands of Panay,
Negros, Cebu and possibly Masbate listed a Vulnerable. The small population
(approximately 100) on Japan's 710 km² Tsushima Island, considered the same
subspecies as occurs in northeastern mainland Asia, has decreased over the last
30-40 years (Izawa et al. 2007).
Population Trend: Stable
Ecology and behavior
Leopard cats are solitary, except during breeding season. Some are active during the day, but most hunt at night, preferring to stalk murids, tree shrews and hares. They are agile climbers and quite arboreal in their habits. They rest in trees, but also hide in dense thorny undergrowth on the ground. In the oil palm plantations of Sabah, they have been observed up to 4 m (13 ft) above ground hunting rodents and beetles. In this habitat, males had larger home ranges than females, averaging 3.5 km2 (1.4 sq mi) and 2.1 km2 (0.81 sq mi) respectively. Each male's range overlapped one or more female ranges. Leopard cats can swim, but seldom do so. They produce a similar range of vocalisations to the domestic cat. Both sexes scent mark their territory by spraying urine, leaving feces in exposed locations, head rubbing, and scratching.
The species can range up to 3,000 m in parts of its range,
which extends into the Himalayas along river valleys. It occurs in a broad
spectrum of habitats, from tropical rainforest to temperate broadleaf and,
marginally, coniferous forest, as well as shrub forest and successional
grasslands. The northern boundaries of its range are limited by snow cover; the
leopard cat avoids areas where snow is more than 10 cm deep. It is not found in
the cold steppe grasslands, and generally does not occur in arid zones, although
there are a few records from relatively dry and treeless areas in Pakistan.
Leopard cats occur commonly in dense secondary growth, including logged areas,
and have been found in agricultural and forest (rubber tree, oil palm,
sugarcane) plantations. The species can live close to rural settlements. Leopard
cats are excellent swimmers, and have successfully colonized offshore islands
throughout their range (Nowell and Jackson 1996; Sunquist and Sunquist 2002). In
the most comprehensive study, based on a large sample size of 20 radio-collared
cats in Thailand's Phu Khieu Wildlife Sanctuary, mean home range size
(95% MCP) was 12.7 km², larger than in other areas of Thailand (4.5 km²)
(Grassman et al. 2005), on Borneo (3.5 km²: Rajaratnam 2000), or on Japan's
Iriomote island (Schmidt et al. 2003). There was no significant difference
between male and female home range size. Open and closed forest habitats were
used in proportion to their occurrence, and activity patterns showed crepuscular
and nocturnal peaks. On Borneo, Rajaratnam et al. (2007) found that leopard cats
hunted rodents in oil palm plantations, and used forest fragments for resting
and breeding. Murids dominate the diet (85-90%: Grassman et al. 2005b,
Rajaratnam et al. 2007). Other small mammals, eels and fish have also been
reported, as well as occasional scavenging of carrion (Nowell and Jackson 1996).
Leopard cats are carnivorous, feeding on a variety of small prey including mammals, lizards, amphibians, birds and insects. In most parts of their range, small rodents such as rats and mice form the major part of their diet, which is often supplemented with grass, eggs, poultry, and aquatic prey. They are active hunters, dispatching their prey with a rapid pounce and bite. Unlike many other small cats, they do not "play" with their food, maintaining a tight grip with their claws until the animal is dead. This may be related to the relatively high proportion of birds in their diet, which are more likely to escape when released than are rodents.
Reproduction and development
There is no fixed breeding period in the southern part of its range; in the colder northern range they tend to breed around March or April, when the weather is mild enough to support newborn kittens. The estrus period lasts for 5–9 days. If the kittens do not survive, the female may come into heat again and have another litter that year. After a gestation period of 60–70 days, two to four kittens are born in a den, where they remain until they are a month old. The kittens weigh about 75 to 130 grams (2.6 to 4.6 oz) at birth and usually double their weight by age of two weeks; at five weeks, they are four times their birth weight. The eyes open at ten days, and the kittens start to eat solid food at 23 days. At the age of four weeks, the permanent canines appear, and the kittens begin to eat solid food. Leopard cats have lived for up to thirteen years in captivity. Leopard cats usually pair for life and raise their kittens together for about 7 to 10 months. Full maturity is reached at 18 months, but in captivity, the male can become ready to breed at 7 months, and the female at 10 months.
In China, leopard cats are hunted mainly for their fur. Between 1984 and 1989, about 200,000 skins were exported yearly. A survey carried out in 1989 among major fur traders revealed more than 800,000 skins on stock. Since the European Union imposed an import ban in 1988, Japan has become the main buyer, and imported 50,000 skins in 1989. Although commercial trade is much reduced, the species continues to be hunted throughout most of its range for fur, for food, and as pets. They are also widely viewed as poultry pests and killed in retribution. In Myanmar, 483 body parts of at least 443 individuals were observed in four markets surveyed between 1991 and 2006. Numbers were significantly larger than non-threatened species. Three of the surveyed markets are situated on international borders with China and Thailand, and cater to international buyers, although the leopard cat is completely protected under Myanmar's national legislation. Effective implementation and enforcement of CITES is considered inadequate.
China, the centre of its range, commercial exploitation has
been heavy: hundreds of thousands of Leopard Cat skins per year were exported in
the 1980s. Although commercial trade is much reduced, the species continues to
be hunted throughout most of its range for fur, for food, and as pets. They are
also widely viewed as poultry pests and killed in retribution. Island
populations are small and seriously threatened in the Philippines and Japan.
Leopard cats can hybridize with domestic cats, as is shown by the popular
domestic breed, the "safari cat". Hybridization in the wild has been reported,
but is not considered a significant threat. Although the species is less
dependent on forest cover than others, habitat loss and fragmentation is still a
major threat across most of its Asian range (Nowell and Jackson 1996).
Included on CITES Appendix II; populations in Bangladesh, India and Thailand are included on Appendix I (as Prionailurus bengalensis bengalensis). The species is protected at the national level over part of its range, with hunting prohibited in Bangladesh, Cambodia, Hong Kong, India, Indonesia, Japan, Malaysia, Myanmar, Nepal, Pakistan, Philippines, Russia, Thailand and Taiwan, and hunting and trade regulations in place in South Korea, Lao PDR and Singapore (Nowell and Jackson 1996, A. Wilting pers. comm. 2008). The species is on Afghanistan’s 2009 Protected Species List, banning all hunting and trading of this species within the country. It is found in numerous protected areas.